Frederick S.

Szalay
Department of Anthropology, Hunter College, CUNY, 695 Park Avenue, New York, N.Y. 10021, U.S.A.

The Foot of Assessment

Oreopithecus: an

Evolutionary

John H. Langdon
Department of Biology, Universityof Indianapolis, 1400 E. Hanna Ave, Indianapolis, IN 46227, U.S.A.
Received 1 May 1986 Revision received 12 March 1987 and accepted 18 March 1987 Publication date August 1987

Keywords: Oreopithecus,foot
morphology and function, Miocene hominoids, catarrhine relationships.

A nearly complete composite foot of Oreopithecusbambolii is described and analyzed both mechanically and comparatively. Unlike the probable primitive anthropoid and catarrhine conditions in which the tarsus was considerably elongated, as in monkeys, for efficient leaping, this region is dramatically foreshortened in the T u s c a n catarrhine. T h e phyletic reduction in the length of the tarsus exceeds that seen in the Miocene sample of Proconsul footbones. Although the foot of Oreopithecusis most similar in its inferred mechanical abilities to living chimps, the details of morphology and subtle but distinct morphological solutions to expedite virtually identical function (mechanics) strongly contradicts the hypothesis that the similarities are homologous to the shared pedal similarities observed in the living apes. The known pedal morphology of parapithecids, monkeys and hominoids is discussed in order to place Oreopithecus in a meaningful phylogenetic and functional perspective. Oreopithecus, with its decidedly advanced catarrhine pedal complex for climbing behavior, shows several modifications, namely: general robusticity of the tarsals and metatarsals; an exceptionally short load arm on the lever system of the calcaneus; strong flexor fibularis; pronounced plantar process on the calcaneal tuber; lack of tight upper ankle joint constraints; mediolateral as opposed to dorsoplantar movements favored by the astragalar head. These form-function attributes of the foot are different from those correlated with habitual pedal hanging in Pongo. The small astragalar head and the relatively undeviated long neck in the Asian ape are in revealing contrast to the conditions in Oreopithecus, although the 1958 skeleton, an adult male, was probably as large as a fully developed female orang, in the 30-35 kg, or heavier, size range.

Journal of Human Evolution (1986) 15, 585 621

Introduction The first report of a few elements of the tarsus of Oreopithecus was given by S traus (1963) in his account of the whole skeleton. More than ten years after the actual time of study by Straus one of us (FSS) spent two weeks in Basel in 1972 where, due to the courtesy of D r J . Hfirzeler, he was able to study the bones reported on as well as additional elements of the foot, an area missing from the 1958 skeleton. Szalay also studied the latter specimen in Florence during that year and some of this information gained from the postcranium was included in the account of Oreopithecus in Szalay & Delson (1979). As a result of the revival of interest in Oreopithecus and because of a recent in depth review by one of us (JHL) of the Miocene hominoid and other catarrhine pedal evidence, we jointly restudied casts and stereophotographs of the unique pedal material. The account below concentrates on the specimens of the foot (all shown on the stereophotos) of which Basel no. 36 presumably represents elements of the left and right foot of the same animal. According to Dr Hfirzeler, all the bones catalogued Basel no. 36 were taken out of the same block of coal. This block was cast prior to its disassembly and this cast is available for inspection in the Basel Natural History Museum. We may now briefly state the aims of our study as follows: (1) to document and selectively compare the rare specimens of the tarsus and pedal cheiridia of Oreopithecus; (2) to decipher the meaning of morphology of these fossils in terms of their mechanics; (3) to associate with the greatest probability possible the form-function attributes of Oreopithecus
0047-2484/86/070585 + 37 $03.00/0 9 1986 Academic Press (London) Limited

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with the biological role (biorole) of the foot of living primates; (4) to suggest phylogenetic relationships of Oreopithecus, based on what we consider the most probable character transformations of aspects of the skull, dentition, and postcranium; and (5) to outline a tentative working hypothesis of catarrhine positional (Prost, 1965) evolution based on the interpretation of the skeletal morphology of fossil and living species.

Definitions of the Concepts "Generalized" and "Specialized" and the Taxonomic Groupings Employed in this Paper
The terms "generalized" and "specialized" are ubiquitous in the literature dealing with the evolution of the positional behavior of catarrhines as well as in the evolutionary literature in general and the literature of the primates in particular. We consider the ambiguous usage of these terms, whose multiple phylogenetic, ecological, and ontogenetic meanings are almost never specified, to be confusing to anyone's attempt to understand what a student is saying about a particular aspect of an animal. These terms are clearly important in the description of functionally or adaptationally related attributes of organisms. We do not advocate their abandonment but we suggest their nonhistorical usage. A structure may be considered generalized if its design enables the animal to perform a number of functions related to a large number of bioroles. A structure may be considered specialized for a function if its design enhances the effectiveness of efficiency of the performance of that function and consequently reduces the effectiveness of other functions related to different bioroles. The language of systematic biology is usually associated with phylogenetic, i.e. genealogical and adaptational, studies. There should be no reason for this as the evolution of diversity is the concern of specialists in ecology, morphology, and behavior as well. Nevertheless the customary designation of antecedent (primitive) and derived (advanced) conditions in accounts of historical sequences in descent have long been used in evolutionary studies as transformation from a "generalized" to a "specialized" state. These terms have further come to designate more diverse and more restricted behaviors, respectively, for ecologists and functional anatomists, and also for many taxonomists. Yet organisms, in spite of prevalent efforts to boost the concept of "living fossils", are neither primitive nor advanced; only their characters may be objectively designated as such. Because any paper dealing with aspects of comparative biology should refer precisely to taxa on several categorical levels, at least within a single study, we find it imperative to define our taxonomic concepts by the genera included in them. The last decade, coincident with the great rise in interest in Miocene higher primates, has seen a great proliferation of family level taxonomic concepts (names) which are neither adequately defined nor diagnosed. We believe that given the nearly universally accepted family level recognition of the Hominidae, all other catarrhines may be accomodated either in the Parapithecidae, Oreopithecidae, Cercopithecidae, Hylobatidae, or the paraphyletic concept of Pongidae. The constantly shifting sand of generic boundries of quite homogeneous species groups (one of the reasons for the experts' disagreements on generic limits) can be easily served and affinities pondered freely by the use of subfamilies, tribes, and subtribes within a broad concept of Pongidae. Until unquivocal delineation of the various clades within a broad concept of the Pongidae becomes possible we see no rationale, evolutionary (phylogenetic), cladistic, or phenetic, for the recognition of new families or superfamilies each time a genus

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587

is j u d g e d to be distinct, when the noted suprageneric categories may still be used. T o posit that the catarrhines, based on the currently available record of diversity, should be subdivided into ten to twelve families (not an u n c o m m o n position taken by many authors who will refer to single genera by family names), compared, for example, with the 9 platyrrhines which are a c c o m m o d a t e d in two families, is perhaps to a b a n d o n any pretense of zoological objectivity for catarrhine classification. T o gain a cladistic classification which is isomorphic with the ephemeral cladograms of various researchers is to reject any reason for a classification which is supposed to aid communication. I f that is the case then why bother with a classification, as the cladograms undoubtedly will continue to represent everyone's working hypothesis of some very tentative notions of splittings--not phylogeny. T h e following concepts within the Pongidae, with due regard given to the priority of the names themselves, will allow most future phylogenetic hypotheses: Ponginae (to include the Gigantopithecini, Ramapithecini, and Pongini); Gorillinae (Pan, Gorilla); Dryopithecinae (Dryopithecini and Proconsulini); Pliopithecinae (Pliopithecus); Propliopithecinae ( Propliopithecus/ Aegyptopithecus ) .

Form-function and Comparative Assessments (Table 1)

Astragalus
T h e astragalus is assessed from two ditt~rently preserved specimens from the left side (Figures 1-9). T h e more complete specimen (Basel no. 36), slightly larger than the other, is crushed in the medial side of the lateral tibial trochlea (ATil). This area is perfectly

Figure 1. A reconstruction of the left tarsus and metatarsus of the foot of Oreopithecus bambolii. The dorsal view is somewhat artificially "flattened" in order to show the relative position of the bones to one another. The hallucial metatarsal is shown in its extreme abducted position. All solid lines shown are known from either the left or right side. Scale represents 2 cm.

588

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SZALAY AND

j. H. LANGDON

preserved in the other specimen. In spite of the damage on one of the specimens, therefore, the pedal conformation of the upper ankle joint (UAJ) is well preserved. The groove for the tendon of the flexor fibularis is pronounced on the underside of the sustentaculum and the prominent lateral tubercle on the proximal astragalus also testifies to a well developed tendon. The distal extension of the astragalar medial tibial facet (ATim) forms a cup shaped articular basin, as in the most African hominoids, not as in Pongo, but also not as in the terrestrial cercopithecines such as Papio. The lack of this cup in forms like Cebus or Pongo, for example, which clearly differ from Oreopithecusin this regard, indicates free and continuous proximo-distal mobility between the astragalus and tibia. The lateral crest of the trochlea is straight and sharp whereas the medial one, as in living gorillines, is more poorly defined. The trochlear surface of the astragalus in Oreopithecusfollows a pattern characteristic of other catarrhines. The lateral crest is relatively higher and longer than the medial crest. The medial crest flares outward (medially) on the astragalar neck to create an articular basin, noted above, for horizontal support of the tibial malleolus. This contributes to the "wedging" of the troehlea, causing the astragalotibial surface to widen distally. The development of this articular basin, the sharpness of the crests, and the form of the AFi (astragalofibular) facets all relate to the degree of accessory movement in all degrees of freedom in the UAJ.

Table 1

The following abbreviations are used in the text and figures in connection with the lower leg and foot. These are similar, utilizing the same approach to those found in Szalay (1982, 1984, 1985) and Dagosto (1985). Specific joints are abbreviated by the combination of the first letters in capitals of the names of those units which make up the joint, and the letter "J" for Joint. Abbreviations entirely in lower case designate bone processes, ligaments, tuberosities, grooves, tendons, etc., or anatomical directions
astragalofibular astragalonavicular astragalonavicular joint astragalus anterior plantar tubercle calcaneus calcaneoastragalar distal calcaneoastragalar proximal calcaneoastragalar calcaneocuboid calcaneofibular calcaneocuboid joint calcaneonavicular joint calcaneonavicular ligament cuboid distal ectocuneiform entocuneiform-metatarsal I joint entocuneiform fibular lateral LAJ m Mc mm Mt Na NCJ NCu ne NEc NEn NMe p pp sa sc Su Sud Sup tca UAJ lower ankle joint medial mesocuneiform medial malleolus metatarsal navicular naviculocuboid joint naviculocuboid neck of astragalus naviculoectocuneiform naviculoentocuneiform naviculomesocuneiform proximal peroneal process sulcus astragali sulcus calcanei sustentacular distal sustentacular proximal sustentacular tuber of calcaneus upper ankle joint

AFi AN ANJ As at CA CaA CaAd CaAp CaCu CaFi

ccj CNJ
cnl Cu d Ec EMT1 En Fi

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589

Figure 2. Oreopithecus bambolii, stereophotos of left, dorsally damaged astragalus and calcaneus (both no. 37). From the top down: dorsal, plantar, lateral, and medial views, respectively.

~!I! ~~

~ii~i~i~i

~i ~ ....

590

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AND

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~Figure 3. Oreopithecus bambolii,stereophotos of left astragalus and calcaneus (both no. 37). Top: distal viewofcalcaneus (above) and astragalus (below); bottom: proximal view ofastragalus (above) and calcaneus (below).

'IL /P
The significance of the wedging of the trochlea, discussed by Langdon (1986), is still poorly understood, yet we believe it to be essentially related to the degree of conjoint adduction in the UAJ when the foot is plantarflexed, and/or inverted. To express this in another way, the tilt of the medial end of the UAJ axis upwards and forwards is accommodated by the elimination of the proximomedial corner of the trochlea, giving it a wedged appearance. Because the fibula rotates and the astragalus adducts in plantarflexion, the joint does not loosen as much as many other authors have assumed. While wedging may be related to plantarflexion or to a more inverted carriage of the foot (particularly conjoining inversion), the exact functional implications of varying degrees of wedging and asymmetry which can be seen among living catarrhines is not clear. In Oreopithecus the extent of wedging compared to Apidium, for example, is relatively great. Oreopithecus

FOOT OF O R E O P I T H E C U S

59]

exhibits a m o d e r a t e l y high l a t e r a l crest, relatively h i g h e r t h a n hominoids which it resembles functionally, yet lower than t h a t of cercopithecids (see Figures 6, 10 a n d the index o f trochlear a s y m m e t r y in T a b l e 2). T h e trochlea of Oreopithecus is tlattish both from side to side (see index of height o f t r o c h l e a r arch in T a b l e 3) a n d front to b a c k (see index of t r o c h l e a r d e p t h in T a b l e 3).

Figure 4. Oreopithecusbambolii, stereophotos of left astragalus (no number) and the left Mt 4 and right Mt 5 (both no. 36). Top: from left to right Mt 5, Mt 4, medial views; astragalus, dorsal view. Below: from left to right: lateral view of metatarsal and plantar view of astragalus. Both in the top and bottom stereos the broken piece on the top right represents the fragment of the astragalar CaA facet.

tl
a

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LANGDON

-Figure 5. Oreopithecus.bambotii, stereophotos of left astragalus (no number) and Mt 4 and 5 (both no. 36). Top: from left to right Mt 5 and Mt 4, dorsal views; astragalus, medial view. Bottom: from left to right: Mt 5 and Mt 4, plantar view; astragalus, lateral view.

I n c a t a r r h i n e s the A F i facet usually flexes laterally at a line c o r r e s p o n d i n g a p p r o x i m a t e l y to the chord of the arc of the lateral crest. This line o f flexion is a s h a r p l y defined crease in the cercopithecids, b u t forms a more g r a d u a l concavity a m o n g hominoids. I n Oreopithecus, the line o f flexion lies more to the vertical so that the a n t e r i o r p o r t i o n of the facet swings laterally. T h e j o i n t m u s t have been loosely fitted; otherwise this would have interfered with the a b d u c t i o n of the foot which usually a c c o m p a n i e s dorsitlexion. Except in possibly an e x t r e m e plantarflexed position, the fibula w o u l d have no significant horizontal s u p p o r t for t r a n s m i t t i n g b o d y weight.

Figure 6. Oreopithecus bambolii, drawings, slightly reconstructed of left calcaneus [(a)-(e)] and left astragalus [(f)-(j)], based on the astragali and calcaneus shown above. Dorsal [(a), (t)], plantar [(c), (h)], lateral [(b), (g)], medial [(d), (i)], and distal [(e), (j)] views, respectively.

(o)

(c)

(f)

(h)

ib)

d)

c j)

i)

4cm
I I I I I

ijl

.yl D .Q

Figure 7. Comparative overview of the dorsal aspect of the astragalus in Cebus (a), a Fayum parapithecid (b), Cercopithecus aethiops (c), S Papio hamadrayas (d), Hylobates sp. (e), Pliopithecus (f), Proconsul africanus, R U 2036 (g), Pan troglodytes (h), Pongo pygmaeus (i), and Oreopithecus (j). Scales represent 1 cm. Note the arrows pointing to the lateral proximal process, area of attachment of one of the strong 0 calcaneoastragalar ligaments. This is partly reconstructed in Oreopithecus due to some damage to this area on the known specimens. T h e relative

0 deviation of the astragalar neck is a good predictor of the importance of grasping for climbing and the degree of cursoriality in these primates, 0 except for Pongo. The undeviated and long astragalar neck with its small head in the latter is causally correlated with pedal hanging in that 0 species. Contrast this condition with Oreopithecus. For abbreviations see Table 1,

(a)

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FOOT OF

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596

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._v.cJ ~-~

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Figure 10. Comparisons of the articulated astragalus and calcaneus in dorsal (above), lateral (middle), and distal (below) views in Pan troglodytes (a), Pongo pygmaeus (b), Oreopithecus bamDolii (c), Papio hamadrayas (d), and Proconsul africanus (e). Scale represents 1 cm. The dished distal articulation of the tibial malleolus on the astragalus (1) is relatively shallow in Pan and Oreopithecus, and it is virtually nondiscernible in Pongo compared to its prominent expression in Papio and Proconsul. T h e deep cleft (2) on the lateral side of the calcaneal tuber, probably related to atragalocalcaneal ligamentous binding, is exceptionally developed in Oreopithecus and Papio (and in Mesopithecus and Dolichopithecus, not shown here). Note the absence of a plantar heel process (probably a primitive catarrhine feature ) in Papio and other cercopithecids. The extreme lateral rotation of the CaCu facet in Pongo sharply distinguishes this clade from known Gorillinae.

(b) (c)
(e)
e
i

(d)
I

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I"C
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1
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9 9 9 O b~

ocu cocucocu,acu

CoCu

598
Table 2

F. S. SZALAY AND J. n. LANGDON

Dimensions of the Oreopithecusfoot bones with means of comparative sample (in mm). All comparative data from Langdon (1984)

Genus and sample size

Oreopithecus
1-3 Trochlear breadth (TA1) 11-8 Lateral crest length (TA4) > 174, 195 est. Trochlear depth (TA5) 0.7 Trochlear asymmetry (TA6) 1-3 Lateral crest height (TA7) n.a., 5'5 Talar head breadth (TA8) 15-8, 16.2 Length of head and neck (TA9) 19"4, 18'7 Calcaneal length (CA1) 38.5 Calcaneal tuberosity length (CA2) 16.3 Posterior talar facet length on calcaneus (CA6) 14'7 Posterior talar facet breadth on calcaneus (CA7) 13'2 Calcaneal tuberosity height (CAB) 21.6 Cuboid facet breadth on calcaneus (CA10) 18.0 Navicular thickness, anterior-posteriorly (NA2) 8"0 Talar facet length on navicular (NA3) 18"2 Talar facet height on navicular (NA4) 12"1 Cuboid facet length on navicular (NA5) 6.2 Cuboid facet breadth on navicular (NA6) 5-2 Navicular facet length on entocuneiform(MC1) 13.5, 12'8, 12"6 Entocuneiformlength (MC2) 13'3, 11.9, 13"7

Papio
18 15-16 20.41 1"34 2"42 7.74 16.94 20"31 50-17 19"62 14.16 12.78 20"33 16.56 9.87 17'72 12.30 7"69 7"66 9"60 15.54

Pan
21 18.04 25.77 0.91 0.32 14-61 23.05 25.55 53.84 19.47 22.02 17.03 31.27 21.48 9.55 23.63 16.51 10.40 6.14 17.45 20.83

Pongo
22 19.08 23.58 1.75 2.61 7-10 19.29 27.91 52.73 17-96 21.08 16.65 29.98 20-40 8.60 21-04 14.72 11.69 7.73 12.89 17.80

T h e short neck (see indices of T A H E A D a n d T A N E C K in T a b l e 3) is more divergent in

Oreopithecus than perhaps in a n y other catarrhine, with obvious implications for grasping ability. T h e head and the a s t r a g a l o n a v i c u l a r (AN) facet are extremely similar to that of Pan
both in shape a n d orientation. T h e size a n d a n g u l a t i o n of the head are such that they were u n d o u b t e d l y capable of w i t h s t a n d i n g considerable forces. T h e relatively large astragalar head must be emphasized here because it is an i m p o r t a n t diagnostic feature distinguishing the advanced climbers like hylobatids or African apes. I n hominids the large head size is due to different functional requirements: T h e significance of the a n g u l a t i o n of the neck with the large head lies in the direction of the compressive forces it can best absorb. This is well to the medial side, as one would expect from a grasping foot in a weight b e a r i n g configuration.

Calcaneus
T h e p l a n t a r side of the astragalus makes up the astragalar half of the lower ankle j o i n t (LAJ). Its features are treated with the conarticular surfaces of the calcaneus. T h e latter is k n o w n from a single specimen (Basel no. 36) on which there is slightly d o w n w a r d crushing on the calcaneal calcaneoastragalar (CaA) facet. T h e deep t u b e r has a well developed p l a n t a r process (pp), implying, as discussed in detail by S a r m i e n t o (1983), that the origin of the superficial head of the flexor digitorum brevis a n d also p r o b a b l y a p l a n t a r

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Table 3

Selected indices with means of comparative samples. All comparative data from Langdon (1984)

Genus and sample size

Oreopithecus
1-3 Index of height of trochlear arch (TAARCHT) Index oftroehlear asymmetry (TAASYM) Index oftroehlear depth (TADEPTH Index oftalar head breadth (TAHEAD) Index oftalar neck length (TAN ECK) Navicular robusticity (NAVROB) Index of ento-cuneiform robusticity (MCHT) Index of length of posterior facet on calcaneus (CA6/CA) Index of breadth of posterior facet on ealcaneus (CA7/CA 1) Index of calcaneal tuberosity length (CACTLEN) Index ofcalcaneal tuberosity height (CACTHT) Index of breadth ofcuboid facet on calcaneus (CA10/CA1) Index of shape ofcuboid facet on navicular (NAVCF) -<28 11 5"9 134, 129 100 379 102, 108, 92 38 34 42 56 47 84

Papio
18 37.9 16"1 8-9 112-6 100"0 304'6 61"9 28'0 25'5 39.1 38'3 33-0 103.9

Pan
21 34"3 1"8 5' 1 128'3 99"8 424"7 84-0 40-9 31.6 37.4 50"9 39.9 59-6

Pongo
22 30.1 9'3 9"3 101-2 120"2 388'0 73'3 40'0 31"4 33"8 50"9 38"9 66-2

a p o n e u r o s i s from this process s t r o n g l y predicts powerful g r a s p i n g and slow c l i m b i n g in which h a n g i n g can be a c o m p o n e n t (see index of caleaneat t u b e r o s i t y height in T a b l e 3). T h e c a l c a n e a l t u b e r also angles p l a n t o m e d i a l l y , p e r m i t t i n g the triceps surae to c o n t r i b u t e to inversion. I n a d d i t i o n to the d e e p a n d r o b u s t construction, a m o d e r a t e heel length limits b e n d i n g stresses a n d would p r o v i d e a m p l e p o w e r for a short load a r m to the m i d t a r s a l b r e a k in the foot or to the base of the h a l l u x (see index of calcaneal tuberosity length in T a b l e 3). A d i a g n o s t i c feature of s o m e w h a t obscure m e c h a n i c a l significance is the exceptionally s h a r p crest r u n n i n g dorsally from the C a A facet to the d o r s a l tip o f the tuber. Similarly, a very extensive a n d d e e p pit on the lateral side o f the tuber, for the l i g a m e n t o u s connection between this area a n d the p l a n t o l a t e r a l process on p r o x i m a l end of the a s t r a g a l u s occurs to this e x t a n t only in Mesopithecus a m o n g the catarrhines. Distally the calcaneus of Oreopithecus is exceptionally shallow, this b e i n g p a r t l y the result of a very small anterior p l a n t a r tubercle for the a t t a c h m e n t of the c a l c a n e o c u b o i d ligment. T h e m e c h a n i c a l significance of this, briefly a l l u d e d to by S a r m i e n t o (1983), m a y be that in climbers increased m o b i l i t y in the tarsus is needed, w h e r e a s in a terrestrial form (e.g., Papio has an e n o r m o u s process for the a t t a c h m e n t of the c a l c a n e o c u b o i d ligament) p r a c t i c i n g bursts of speed in leaping a n d running, a r o b u s t l y b r a c e d calcaneus and cuboid is a d v a n t a g e o u s . I n any case, the i m p o r t a n c e of this trait is in connection with the b r a c i n g of these two bones a n d only indirectly with h a b i t a t preference. A large a n t e r i o r p l a n t a r tubercle can be as a d v a n t a g e o u s in an a r b o r e a l l e a p e r as in a terrestrial runner.

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As noted above, the load arm of the calcaneus in Oreopithecusis exceptionally short, in keeping with the general brevity of the entire tarsus. This short load arm of the calcaneus accentuates the extreme lateral spread of the LAJ. The CaA facet is relatively long as in pongids (see indices of length and breadth of the posterior facet on the calcaneus in Table 3), although this is obscured by its great breadth. The great breadth was in fact responsible for the erroneous assessment by one of us (Szalay, 1975) that this facet was short and therefore cercopithecid-like. The greater length of the calcaneal CaA facet compared to its astragalar counterpart, unlike in cercopithecid monkeys and hominids, clearly indicates extensive LAJ movements. The distally broader CaA facets, unlike those in Pongo where they are proximally broader, indicates the well developed ability of the foot to abduct in the LAJ, an important attribute of climbers. The relatively great surface area relates both to phylogenetic increase in excursion from a more restricted ancestry and to body size in the pongids. The necessity for greater mediolateral accommodation proximally in the LAJ, as in orangutans (Langdon, 1986) and in advanced tree squirrels (Szalay, 1985), reflects the movements necessary when the animal is suspended by its foot. The sustentaculum is well developed and gently concave to support the talar head and neck throughout excursion. Such a design, well displayed in the living pongids, indicates both mobility and secure close-packing of the astragalar head in the weight supporting inverted position. Cercopithecids, by contrast, greatly restrict mobility with a short calcaneal CaA facet, as noted above. Their narrow and waisted sustentaculum, however, is shared both with platyrrhines and most strepsirrhines which have a long and helical CaA facet and subsequently a highly "invertable" LAJ (see Figures 8, 9). In spite of the similarity of the sustentacular morphology of cercopithecids to taxa noted, there are some significant differences between them. In the cercopithecids the long sustentaculum is often disconnected and the two halves are always set obliquely to one another, mirroring a greater restriction of movement in the LAJ than in the strepsirhines or platyrrhines. In fact these sustentacular units in cercopithecids contact different subfacets of the astragalar head and firmly embrace the latter. This condition in Old world monkeys which virtually restricts the invertability of the LAJ is a highly derived condition among the anthropoids. In contrast, it is the relative flatness of the sustentacular facet in hominoids and Oreopithecus which permits extensive motion in the LAJ of these forms. Although in platyrrhines this facet is not flat, its smooth curvature permits ample LAJ mobility. It may be added here that the restriction displayed by cercopithecids appears to be an important aspect of the stabilizing mechanism of the LAJ during high speed running on level substrate.

Cuboid
The cuboid is known from a right specimen which is undamaged (Basel no. 35, Figures 11-14). This bone of Oreopithecus, as that of the Miocene hominoids (Langdon, 1986) and living pongids, is characterized by a deep pivot-like CaCu facet, which allowed helical rotation of the foot on the calcaneus. Quite distinct from the hominoids, however, the cuboid of Oreopithecusis dorsoplantarly flattened. Although this is superficially ateline-like, we believe that this flattened condition is independently evolved from an unknown, perhaps Cebus or Aotus-like condition of the anthropoid morphotype. The calcaneocuboid joint (CCJ) is broad and moderately deep (see index of breadth of cuboid facet on calcaneus in Table 3). The dorsolateral portion of the articulation slopes posteriorly to form a sellar surface. Living and Miocene pongids typically flatten and

FOOT OF OREOPITHECUS

601

Figure 11. Oreopithecus bambolii, right entocuneiform and cuboid (both no. 36). From top to bottom: entocuneiform in plantolateral, medial, dorsal, distoplanar, and distal views, respectively; cuboid in plantar, dorsal, lateral, medial, and distal views, respectively.

602

F.S.

SZALAY

AND

j.

H,

LANGDON

Figure 12. Oreopithecusbambolii, two left entocuneiforms, terminal pedal phalange, proximal half of left Mt 3, proximal two thirds of right Mt 1, and curved pedal phalange. All specimens are no. 36, except for the entocuneiform labelled no. 65. Entocuneiforms in medial (top) and proximal (bottom) views; terminal and intermediate phalanges in side (above) and plantar (below) views; Mt 3 in dorsal (above) and lateral (below) views; Mt 1 in lateral (above) and plantar (below) views.

1 1
extend this region to enhance distribution of forces, while in cercopithecids and platyrrhines such as Saimiri, Cebus, and possibly the anthropoid morphotype, the form of the calcaneal C a C u facet resembles a uniform conical concavity. I n Oreopithecus the calcaneal calcaneocuboid (CaCu) facet with its deep depression and dorsolaterally facing side allowed exceptional mediolateral mobility of the foot and dorsiflexion on the calcaneus. This facet is shallow dorsoplantarly, somewhat Pan-like, but its overall shape is unique. T h e facet does not show any sign of the medial displacement displayed in Pongo. T h e dorsoplantar shortness of the calcaneal and cuboid C a C u facets strongly suggest significant midtarsal flexion with insufficient support for the rigid lever action required by both cercopithecids and living hominoids. T h e groove on the plantar surface of the cuboid for the peroneus longus is deep, as in most primates with a strongly adductable hallux. It should be noted here, however, that a large groove for a large tendon of the peroneus longus is prevalent on the cuboids of sundry

FOOT OF OREOPITHECUS

603

Fig. 13. Oreopithecus bambolii, right cuboid, no. 36. Plantar (above left), dorsal (above right), lateral (middle left), medial (middle right), proximal (bottom left), and distal (bottom right) views, respectively. Scale represents 1 ham.

604

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AND

J.

H.

LANGDON

Figure 14. Comparisons of the right cuboid in plantar viewof Cebuscapucinus (a), Papiohamadrayas(b), Pongopygmaeus(c), Pan troglodytes(d), Gorillagorilla (e), and Oreopithecus bambolii (t). Scale represents 1 mm. Note that Oreopithecusshows no special similarity to living pongids.
(a)

(b)

(c)

(d)

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m a m m a l s without a mobile or hypertrophied hallux. T h e peroneal tubercle of Oreopithecus, to which its retinaculum attaches, is very weakly expressed, contrary to predictions based on living catarrhines. A feature of the relatively obscure significance is the conformation of the articular facets for the fourth and fifth metatarsals on the cuboid. In sharp contrast to Pan, for example, the distal surface of the cuboid in Oreopithecus is relatively level (see Figures 13, 14). I n the living great apes the base of the Mt4 has an ovoid facet, whereas the base of the Mt5 has a nearly flat contact with the cuboid. I n cercopithecids and Homo both metatarsals have a nearly flat cuboid contact. As far as we can determine from the single available fossil of Oreopithecus, it is more similar to the condition displayed by some platyrrhines like atelines (or archaic catarrhines?) than to that seen in pongids. This subtle aspect of the cuboid m a y relate to forefoot mobility, more precisely to the tarso metatarsal flexion and extension. In general this is contrary to predictions for Oreopithecus (namely that the foot is exceptionally mobile), but we suspect that the shortness of the metatarsals m a y have played a part in the development of this feature. In hominids this feature clearly relates to the rigidity necessary for running. I n an arboreal form the ability of the foot to curl around a b r a n c h is of p a r a m o u n t importance. T h e longer the foot, the more bending is needed. In a short footed animal like Oreopithecus, however, the need for this mobility at the cuboid-metatarsal

FOOT OF

OREOPITHECUS

605

contact is considerably reduced. We can think of no other way at present to ascertain the mechanical significance of this for the forefoot. Consequently even less may be said of its relevance to the biorole of the whole foot. We are not at all certain whether this feature is a primitive retention from a protocatarrhine ancestry, or merely a functionally, but not necessarily in its biorole, convergent attribute to those of atelines.

Navicular
The navicular is known by one left undamaged specimen (Basel no. 36, Figures 15-17). This bone, like its homolog in living pongids, is proximodistally very shallow, and contrasts sharply to the primitively longer condition in cercopithecids, Miocene hominoids, parapithecids, platyrrhines, and Eocene strepsirhines and haplorhines. The three cuneiform facets arch across the distal border of the navicular rather than sharing the same plane. This pattern is like that found in apes. The distribution of these cuneiform facets suggests great mobility and an orientation for more medially directed forces, at least from the Mt 1, transferred through the entocuneiform. The relative strengthening of these joints which is expressed differently among other catarrhines may be related to the greater stresses generated by the elongated metatarsals. One of the most significant patterns which has emerged from Langdon's (1986) study of the Miocene hominoids was the alignment of the tarsal bones either for more longitudinally or more transversely oriented stresses in cercopithecids and hominoids, respectively. The navicular is highly expressive of this pattern in its arc and robusticity (i.e., its relative foreshortening). Oreopithecusdefinitely fits the grasping pattern displayed by the hominoids. In spite of the extreme and pongid-like shallowness of the navicular, which is a mechanically significant attribute necessary for a habitual climber whose tarsals will be required to flex on each other, there are some significant features which suggest the entirely independent attainment of this condition in Oreopithecus compared to living pongids and Proconsul. In the apes and Miocene hominoids the navieulobucoid (CuNa) facet is triangular; in Pan [Figure 17(c)] this facet is offset plantad from the astragalonavicular joints (ANJ). this long triangular contact in hominoids (see Figure 17) prevents the navicular and cuboid from rolling on one another in a transverse plane. In Oreopithecusthe CuNa facet is rectangular. A hypothesis of homology for the shallowness of this bone is therefore contradicted by the differing details in the two groups. In the living apes and Miocene hominoids the naviculoentocuneiform (NEn) facet is attenuated and decidedly sellar. In Oreopithecus, in spite of the striking adaptive resemblance of the greatly shortened tarsus to that of Pan, this contact is merely hinged. Oreopithecus also differs from the hominoids in having a relatively transversely very short NCu contact and in having an additional process planted to the tubercle for the tibialis posterior. In spite of the general similarities of the navicular of Oreopithecus to hominoids (which display a great adaptive diversity) there is a primitive catarrhine, "monkey"-like (i.e., found in forms like Cercopithecus, Cebus, and others) feature, the non-sellar NEn contact (Figure 18). Given the great range of mobility of the hallux, as reflected by the entocuneiform, discussed below, this difference is a significant support for a hypothesis that Oreopithecus is not a derivative of the pedally known dryopithecine stock.
CurteijCorms

Both left and right entocuneiforms are known, almost certainly from the same animal (Basel no. 36, Figures 11, 12, 19-21). In Oreopithecus, unlike in platyrrhines or

606

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S Z A L A Y AND J.

H. L A N G D O N

Figure 15. Oreopithecus bambolii, stcreophotos, in five views, of the left navicular, left mesocuneiform, and right ectocuneiform (all no. 36). From top to bottom: navicular in proximal, distal, dorsomedial, plantar, and dorsal views; ectocuneiform in proximal, distal, lateral, medial, and dorsal views; entocuneiform in proximal, distal, medial, lateral, and dorsal views.

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FOOT OF OREOPITHECUS

607

cercopithecids, the entocuneiform is proximodistally shortened compared to its dorsoplantar height. This shortening (or increased robusticity) correlates with that of the navicular noted above and signals an increase in the compressive and other forces transmitted from the base of the hallux through to the body of the astragalus. As noted before under the navicular, this is highly characteristic of the advanced and seldom leaping, climbing graspers. These proportions of the bone are probably one of the most extreme among the primates.
Figure 16. Oreopithecus bambolii, left navicular (no. 36), shown in proximal (above right), dorsomedial (bottom left), and plantar (bottom right) views, respectively. Scale represents 1 mm.

Table 4 presents some data on the entocuneiform. This and the morphological information suggest that Oreopithecus paralleled the apes in both the proximodistal shortening and hypertrophy of the proximal facet. Both of these features are causally associated with forces generated by a powerful hallux. Among the living apes Pongo has the longest entocuneiform, reminiscent in its proportions of some of the Miocene hominoid specimens (RU 5872, 2036), cercopithecids, and platyrrhines. Given the proximodistally shortened navicular in the great apes and Oreopithecus, the size (dorsoplantar length) of the NEn facet relative to that of the EnMtl contact is much smaller in Oreopithecusthan in the living apes. In other words, in spite of its greatest length reduction, the proportions of the NEn contact in Oreopithecus are not like the relatively hypertrophied form in the African apes, but like the proportions on the long entocuneiforms of other primates. This attribute

Figure 17. Comparisons of left naviculars in plantar (above), dorsal (middle), and proximal (below) views, respectively, of Oreopithecus bambolii (a), Proconsul africanus, R U 5872 (b), and Pan troglodytes (c). Scales represent 1 mm. Note the adaptively similar shortening of this bone in these taxa, and the sellar nature of the NEn facet (1) shared as a derived feature between Proconsul and Pan. Oreopithecus has an extra bony process (2) not found in other known pongids. For abbreviations see table 1.
0 Co

NEn NEn r 1 Inl

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FOOT OF OREOPITHECUS

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a l o n e s t r o n g l y suggests t h a t the r e d u c t i o n of the l e n g t h o f tarsals in the g r e a t a p e s a n d Oreopithecus w e r e i n d e p e n d e n t p h y l e t i c events. T h e s h a r e d special similarity, a p r o b a b l e s y n a p o m o r p h y , b e t w e e n M i o c e n e h o m i n o i d s a n d living p o n g i d s , o f a slightly sellar N a E n facet on the n a v i c u l a r f u r t h e r s u g g e s t s t h a t the i n d e p e n d e n t r e d u c t i o n o f the n a v i c u l a r l e n g t h in Oreopithecus was n o t f r o m a m o n g the M i o c e n e h o m i n o i d s , at least n o t of the k n o w n d r y o p i t h e c i n e ( a m o n g the p r o c o n s u l i n s ) conditions.

Table 4

Metatarsal d i m e n s i o n s (in ram) and i n d i c e s with m e a n s of comparative samples. All comparative data from L a n g d o n (1984)

Metatarsal II Absolute length (MT1) III -65"97 68"02 91-21 -6'27 8.70 8"88 IV 50.1 67"53 66"36 97"50 7.8 6' 18 7'29 7'93 5"6 6.30 6.41 6"69
100

V 48.6 est. 64.09 58-58 78"40 6-6 6.25 6.10 6.70 6"8 5"27 6-61 7'03
--

Oreopithecus Papio Pan Pongo

Absolute shaft height (MT2)

52'9 62.44 71"35 93"17 8"3 6'01 8.71 8'50

Oreopithecus Papio Pan Pongo

Absolute shaft breadth (MT3)

6'7 -6"19 6.89 6'99 6"80 6.81 6.44 Relative length as percent of MT IV x 100 (MTLEN) Oreopithecus 106 -Papio 92 98 Pan 108 103 Pongo 107 104 Robusticity index (100 x (breadth + height)/length; MTROB) Oreopithecus 28 -Papio 19"8 19.8 Pan 22'2 22"8 Pongo 16"3 16"8 Shaft compression (100 x height/breadth: MTCOMP) Oreopithecus 123 -Papio 97"1 91 "8 Pan 125"0 129"2 Pongo 127"7 140-5

Oreopithecus Papio Pan Pongo

100 100 100 27 18.4 20"7 16"7 139 99"2 114"4 119-8

95 88 89 28 est. 17"9 21"7 17"5 97 119"6 92"7 95"4

T h e m e s o c u n e i f b r m (Basel no. 36, F i g u r e 15), a l t h o u g h u n d a m a g e d , f r o m the left side is p o o r l y u n d e r s t o o d by us. I t is r e l a t i v e l y shorter, c o m p a r e d to the p r o x i m a l tarsals, t h a n those in c e r c o p i t h e c i d s a n d p l a t y r r h i n e s , so in its l e n g t h it is m u c h m o r e as in apes. T h i s is c l e a r l y a g e n e r a l a d a p t i v e s i m i l a r i t y b e t w e e n living a p e s a n d Oreopithecus, a p p l y i n g e q u a l l y to all of the tarsals. T h e n a v i c u l o m e s o c u n e i f o r m ( N M c ) facet is h i g h (see i n d e x o f m e s o c u n e i f o r m r o b u s t i c i t y in T a b l e 3), s i m i l a r to t h a t o f living apes. T h i s is a n o t h e r i n d i c a t i o n t h a t all m e d i a l e l e m e n t s in the foot are well s t r e n g t h e n e d for p o s t e r o l a t e r a l l y d i r e c t e d stresses.

Figure 18. Comparisons of the left navicular in plantar (above), dorsal (rniddle), and proximal (below) views, respectively, of Notharctus tenebrosus (a), Apidium phiomense (b), Cercopithecus aethiops (c), and Cebus capucinus (d). Scales represent 1 mm. Note the great length of the navicular in these forms, very likely a primitive euprimate character reflecting the common grasp leaping ancestry, compared to pongids. The relatively very small NEn facet in Cercopithecus reflects the reduced role of the hallux. The relatively large size of this facet in Apidium suggests the presence of a powerful grasping hallux. For abbreviations see Table 1.

< >

z c; o

FOOT OF O R E O P I T H E C U S

61 l

Figure 19. Oreopithecusbambolii,same specimens as Fig. 12. Entocuneiforms in lateral (middle) and distal (below) views; terminal and intermediate phalanges in side (above and middle) and dorsal (bottom and above) views; Mt 1 in medial and dorsal (both above) views; Mt 3 in ventral (middle), and medial (below) views.

t
Metatarsals and Phalanges

T h e ectocuneiform (Basel no. 36, Figure 15), is known from a right s p e c i m e n ; it is slightly larger d o r s a l l y t h a n the d o r s a l surface o f the mesocuneiform. T a b l e 4 gives its i n d e x of r o b u s t i c i t y c o m p a r e d to some selected catarrhines.

T h e m e t a t a r s a l s (Figures 4, 5, 12, 19), as they are preserved, are unique a n d a d a p t i v e l y r e s e m b l e those in hominoids. T h e second a n d fourth are relatively well preserved, the third

612

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A N D J.

H,

LANGDON

Figtare 20. Oreopithecusbambolii,right entocuneiform, no. 36. Lateral (above left), medial (above right), proximal (middle left), distal (middle right), plantar (bottom left) and dorsal (bottom right), views, respectively. Scale represents 1 ram.

is incomplete, and the fifth is broken in midshaft but rejoined. T h e y are relatively short for the a p p a r e n t size of the animal and very robust (see indices of absolute length and robusticity in T a b l e 4). T h e second metatarsal is longer than the fourth, a characteristic of animals with a strong digital grasp (see index M T L E N in T a b l e 4). The shafts are reinforced in their dorsoplantar diameter (see index of shaft compression in T a b l e 4). T h e robusticity of the shaft of the metatarsals of Oreopithecus is consistent with strong forces from the digital flexors.

FOOT OF OR.EOPITHECUS Figure 21. Comparisons of the right entocuneiform in medial view of Pan (a), Oreopithecus (b), Papio (c), and Cebus. Scale represents 1 mm. Note the great proximodistal shortening of this bone in Oreopithecus. For abbreviations see Table 1.

613

The only intermediate phalanx known (Figures 12, 19) is strongly curved, comparable in that regard to other hominoids. On both sides of the bone the sharp transversely directed flanges suggest powerful digital flexion. T h e single terminal phalanx (Figures 12, 19) is relatively slender and cylindrical, suggesting that it is not a hallucial one which are usually flattened for their special purposes. Its morphology is not unusual for one of lateral digits in catarrhines.
Predictions of Biorole from Some Form-function Attributes

The mechanical capabilities of the specific form-function complexes discussed above may be examined in terms of actual behaviors that the animals with these attributes can perform, the frequency with which they perform them, and the selective importance of different behaviors. The principle of mechanical containment has been proposed and discussed by Langdon (1986). He has suggested that certain behaviors will require form-function solutions which may contain the stress resistance attributes necessary for other postures or movements. This approach, we believe, allows considerable bridging between character complexes otherwise difficult to analyze phylogenetically or adaptively. Persistence of primitive characters alongside the appearance of derived ones allows for a transformational explanation ofbioroles inasmuch as aspects of new behaviors may be seen as being partly contained within the ancestral form-function complex. The principle of containment, therefore, is an essential perspective in a transformational analysis which permits a synthetic (evolutionary) appraisal of morphology for its genealogical, functional, and adaptive significance. The positional behaviors of Oreopithecus were obviously influenced by the body weight of the animal. Judged from all the skeletal remains we believe that the upper end of the weight distribution in Oreopithecus was probably around 35 kg, the size limit shown by an "average" Bornean female orangutan (Napier & Napier, 1967).

614

F.s.

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H. L A N G D O N

In the most general terms the tarsal mechanics of Oreopithecus, with its loose-fitting articulations, suggest climbing as the primary adaptation (clearly all primates climb, but only "climbers" tend to do this at the exclusion of frequent leaping or habitual running). These joints would be inappropriately unstable for rapid loading as occurs in plantarflexion of the foot on the foot, for example, during running and leaping. The large head and the medially deviated short neck strongly suggests vertical climbing as the predominant mode of locomotion, similar to the common ancestor of the Gorillinae, an animal with the climbing habits and abilities of Pan. The form-function of the pedal remains of Oreopithecus, as stressed above, is astonishingly similar, although we believe not homologously so, to that of chimps. The postures, and various locomotor movements practiced with the feet of these animals, therefore, either contain the behaviors of the foot of Oreopithecus, or are basically the same. Clearly we are not saying that the positional behavior of Oreopithecus was the same as we see in Pan. Whatever the total picture of the skeletal morphology of Oreopithecus suggests for the animal, the foot strongly points to a highly evolved climber with a very flexible tarsus and a grasp as powerful as that of any of the similar size Miocene members of the Pongidae. As Sarmiento's (1983) study established for all primates and other arboreal mammals, the plantarly recurved tuber of the calcaneus, a heel or plantar process, is strongly predictive of the powerful toe flexion associated with slow climbing, and, perhaps, with hanging. We found, however, that in mammals which habitually hang (colugos and the orangutan, to mention only a few), the proximal articulation of the sustentacular surface on the astragalus and calcaneus is clearly sellar in shape to allow the necessary adjustment by the hanging body on the firmly anchored foot just above this point. In advanced tree squirrels (Szalay, 1985) the problem of hanging downward on a tree trunk is solved analogously by the transversely widened sustentaculum in the LAJ. The astragalar CaA facet in orangs which habitually practice hanging is widened proximally for the same reasons that the proximal end of the sustentaculum is sellar: it allows additional mediolateral translation in the LAJ and thus relieve stresses that movements of the animal's body may generate. In Oreopithecus, as in chimps, it is the distal end of the astragalar CaAs facet which is widened to allow a greater range of mediolateral movement of the inverted foot, a hallmark of climbers with a great range of movements in the foot. It is significant that in Pongo, which is certainly an able climber, the mechanics associated with climbing alone cannot accommodate or contain the loads generated during habitual hanging, but the severe requirements of hanging the large body in tension will contain the physical necessities of a slow climber. There is no evidence in the foot of Oreopithecusfor pedal hanging to the extent that it is practised in orangs. The great and derived flexibility of the tarsus of Oreopithecus, hallmark of a climbing foot such as that of Pan, is indicative, as in all other hominoids of the Gorillanae and Dryopithecinae, of climbing on irregular supports rather than horizontal travel on large branches or on the ground. These form-function attributes may be briefly enumerated as follows: (a) there is a general robusticity of the tarsals and metatarsals; (b) the lever system ofcalcaneus shows an exceptionally short load arm; (c) there is evidence for a strong flexor fibularis; (d) the digits are relatively very short; (e) Mt2 is (we infer) longer than Mt3; (f) there is a pronounced plantar process of the calcaneal tuber; (g) there were no tight UAJ constraints present as the astragalar trochlea is quite shallow; (h) the head of the astragalus clearly favored mediolateral as opposed to dorsoplantar movements.

F O O T OF

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615

To this list we may add that the size and angulation of the astragalar head gives testimony that the foot was capable of withstanding considerable forces generated by its body weight on a branch grasped by its robust opposable hallux. The range of this opposability is clearly shown on the entocuneiform. In the running-leaping cercopithecids, by contrast, the astragalar head is both more gracile and less deviated to receive the more longitudinally directed stresses. In habitual positions of pedal suspension, the astragalar head and neck are also both more gracile and less deviated to receive the more longitudinally directed tensile stresses. In habitual pedal suspensory behaviors the loads are withstood primarily by the soft tissues and do not need to be resisted by large bony surfaces. The small astragalar head and the relatively undeviated long neck of Pongo are a telling contrast to the morphology seen in Oreopithecus.

Catarrhine Locomotor Evolution and the Relationships of Oreopithecus

Although in its mechanical abilities the foot is most similar to that of a living chimp, it does not necessarily follow that Oreopithecusis also phylogenetically nearest to the living African apes. Let us now briefly examine the known evidence, with emphasis on the foot, from the vantage point of phyletic affinities. We will attempt to present a hypothesis of evolutionary transfbrmation for the form-function attributes of the foot of the known groups ofcatarrhines, a hypothesis which, in our view best accounts for the known details and temporal distribution of the record. No particular genealogy influences our specific ideas of transformation. We would like to note that the path of phyletic descent (i. e., genealogy ) is an abstraction from the fact ofevolu tion of organisms (a total event encompassing considerably more than genealogy) and therefore is dependent on an evolutionary assessment for its various degrees of probability (see Bock, 1977, 1981; Gutman, 1977; Szalay, 1977, 1981a, b; etc. for a non-cladistic approach to phylogenetics). As outlined by Bock (1981), Szalay (1977, 1981a, b, 1985), and many others, it is the plausibility of all known hypotheses for the transformation of known character complexes which is the bioscientifically meaningful approximation of actual taxon phylogeny. This avowedly phylogenetic approach (which seeks to understand hypothesized transformations in order to increase the probability that the assigned stages in the m0rphocline polarity approximate the actual evolutionary events, and not only tally "synapomorphies" generated by implausible methodologies) is different from cladistics which usually omits functional and adaptational character analysis. We believe that because the lack of details of similarity indicate nonhomology between Oreopithecus and the living apes, the form-function similarity of these is independently acquired. The parallel or convergent acquisition of the overall shortening of the tarsals, along with the necessary modifications for climbing and occasional hanging (features like the great mobility of thejoi,nts, the independence of these joints from each other, including the loose construction of the articular relations of footbones) are the features which require careful comparison between Oreopithecus and the pongids. This comparison reveals, as indicated under the examination of form-function, that the similarities between Oreopithecus and any of the extant African pongids is less than the similarity of any of these to one another, or the special similarities between the Miocene proconsulins and the living gorillines.

616

F.S.

SZALAY

AND J.

H.

LANGDON

The most ancient modern primate postcranial evidence, that of the Eocene Adapidae, and less importantly for our discussion, the pedally somewhat anthropoid-like Omomyidae (Szalay & Dagosto, in prep.), display foot attributes which not only show the grasping abilities of the hallux well developed, but also attest to strong commitment to leaping (Szalay & Dagosto, 1980; Szalay & Delson, 1979). This particular locomotor mode, postulated to be an ancestral one for the modern primates (strepsirhine adapids and their descendants) was designated as grasp leaping in order to draw attention to the novel combination of progression in the morphotype. Szalay & Dagosto (1980) have briefly proposed grasp leaping as a morphotype locomotor mode (discussion of the concept in that paper). They hypothesized that the ancient primate grasping adaptation (a feature acquired by the protoprimates or protoarchontans and not a primitive eutherian heritage; see Szalay, 1984) coupled with the development of claws into nails (i.e., claws without sharp points) was utilized in conjunction with leaping when the animal makes use of the grasp as both a take-off and landing strategy among branches. This view was recently strengthened by Dagosto's (1985) study of the upper ankle joint in Paleogene primates. There is nothing contradictory in the behavior of the rapidly running and jumping platyrrhines (the derived modifications of the callithrichine and ateline morphology and behavior notwithstanding). The pedal remains of catarrhines from the Fayum deposits (see descrilotions in Conrov. 1974; Fleagle, 1983; Conroy & Rose, 1983; Fleagle & Simons, 1982) was unequivocally important because of their age and because they clearly indicate two distinct formfunctional approaches to locomotion by early Oligocene times. This dichotomy appears to be between the Parapithecidae and Pongidae (sensu lato), but admittedly the relevant material for the dentally well represented species is still inadequate. Among the parapithecids the crural specializations of Apidium (distal fusion or attenuated juxtaposition) have recently been described (Fleagle & Simons, 1983). In spite of this crural fusion (a similar flattened juxtaposition in some platyrrhines is not uncommon), the relatively great length of the tarsals is shared with primitive adapids and omomyids, and the astragalar and calcaneal morphology of known parapithecids (see Figures 7-9, 22) is most similar to those of Cercopithecus spp., particularly vervets, a species not infrequently found on the ground. This similarity of parapithecids, along with the undoubtedly incipiently bilophodont upper molars of Parapithecus (see Figure 152 G in Szalay & Delson, 1979), to the cercopithecid morphotype may continue to remain a serious impediment to the derivation of the Cercopithecidae from Miocene hominoids, a hypothesis so often evoked during the past decade. In contrast to the parapithecid morphology, discussed in greater detail below, the reported bones of Propliopithecus, and also probably Aegyptopithecus (if the latter concept can be kept distinct from the former) show a telltale sign of strong climbing adaptation in the well developed plantar process (Figure 22). We may predict that once the pes of the Fayum pongids become better known, we will see other indications that the essentially cebine and Aotus like morphology also will show resemblances to that character complex of the Miocene hominoids which suggest ties to hylobatids, Pliopithecus, and the living apes. Langdon (1986) has shown that the Miocene hominoid pedal characters are in many ways "monkey-like" (narrow LAJ, relatively long tarsals, etc). These similarities may be further delineated by stating that they represent ancient primate attributes, some first encountered among Paleogene strepsirhines (Decker & Szalay, 1974). Thus the similarities shared between Miocene dryopithecines like the proconsulins and the cercopithecid

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OREOPITHECUS

617

Figure 22. Comparisons of the medial views of left calcanea, ofApiclium (a), Rusinga Miocene hominoid RU 1654 (b), Proliopithecussp. (c), and Miocene hominoid CA 2270 (d). Scales represent 1 mm. Note the conspicuous absence of the plantar heel process in Apidium compared to the other three forms. All four taxa, along with platyrrhines and cercopithecids share the probably primitively long load arm of the calcaneus, a very likely heritage from either the common anthropoid, haplorhine, or most likely ancestral euprimate of the Paleocene. The similarity of the calcaneus of Propliopithecus to several Miocene pongids provides strong additional suggestion for the evolutionary classification of the Fayum genus within the Pongidae.
(a) (b) (c)

(d)

m o r p h o t y p e are a n t h r o p o i d a n d also c a t a r r h i n e m o r p h o t y p e resemblances. T o derive either of these M i o c e n e groups from the other one would have to reconcile the a p p a r e n t specializations distinct in each group. W h e t h e r the similarities which are derived features in both p a r a p i t h e c i d s a n d cercopithecids are h o m o l o g o u s l y shared is a difficult question given the p a u c i t y of the fossil evidence. I n the form-function of the basic mechanics o f the U A J , LAJ, and A N J , the extremes of j o i n t m o v e m e n t s seen in p l a t y r r h i n e s a n d Miocene h o m i n o i d s are eliminated. T h e configuration of the CCJ, a l t h o u g h s h a r e d between p a r a p i t h e c i d s a n d cercopithecids, i s also present in some p l a t y r r h i n e s a n d cercopithecids certainly indicate a functional f r a m e w o r k which c o n t a i n e d r u n n i n g and leaping. This p a t t e r n u n d o u b t e d l y w o u l d a c c o m o d a t e climbing as well, a l t h o u g h the r e d u c e d ability in the L A J of both g r o u p s to invert the foot signals the selectional d e m a n d s a c c e n t u a t e d which were distinct from their ancestry. W h a t is the evidence which would require us to consider these p a r a p i t h e c i d - c e r c o p i t h e c i d a t t r i b u t e s to be nonhomologous? T h e relatively large N E n facet in p a r a p i t h e c i d s , an a n c e s t r a l e u p r i m a t e feature (see Figure 18), c o m p a r e d to the m o r e a d v a n c e d small facet in cercopithecids, cannot be a serious i m p e d i m e n t to a hypothesis of possible p a r a p i t h e c i d - c e r c o p i t h e c i d ties. T h e shortened tarsus of M i o c e n e hominoids, c o m p a r e d with an hypothesized e l o n g a t e d condition of the c a t a r r h i n e m o r p h o t y p e , along with such features as the relatively large a n d d e v i a t e d a s t r a g a t a r head, the d e e p and helical C C J , tall calcaneal tuber with a heel process on m a n y , a n d the exceptionally developed hallux represents a subtle complex of d e r i v e d

618

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S Z A L A Y AND

J.

H. L A N G D O N

features overlain and mixed with the generally more primitive catarrhine length of the tarsals and the tightly fitting UAJ. These latter features may be indicative of the retention of characters from a more ably leaping ancestry. The persistance of an impressive calcaneal heel process in adult gorillas is certainly instructive in this regard. Even if the heel process has a mechanically meaningful place in gorilla foot function, which it undoubtedly must have, this feature more likely represents a heritage than adaptively innovative attribute for the purposes of ground walking. Rather than eliminate the process and the associated superficial head of the flexor digitorum brevis, causally related to slow climbing and pedal hanging (Sarmiento, 1983), it was retained as part of the total pedal solution (which on the level of the whole pes is an adaptation of that group) of the gorilla habitus. It is highly unlikely that it indicates the strong adaptive commitment of gorillas to those behaviors which shaped the evolution of the heel process. It is important to recognize phylogenetic lag in morphological as well as functional analysis. This is not an attempt to forgo functional explanations but rather to give a theoretically more realistic and complete evolutionary account of adaptively often inexplicable features. Any other theoretical framework for interpreting tbrm and function finds itself creating assumptions which imply that form is fully explicable in functional and biorole contexts. This nonhistorical approach ignores the explanatory perspective of vertical, or transformational, comparisons which place form-function and biorole correlations in an evolutionary, phylogenetic perspective (Szalay, 1981a). To subscribe to the notion that form can be explained by mechanics alone is to deny the existence of phylogeny, and furthermore to eliminate historical explanations of the divergent mechanical solutions to seemingly identical bioroles. A comparison of the calcaneus of Hylobates (Figures 23, 24) and Propliopithecus (Figure 24) with Oreopithecus and other catarrhines reveals the adaptive distinctions between gibbons and Pliopithecus. In hylobatids the bowed calcaneus and the extremely deviated Figure 23. Hylobatessp. Above: dorsal view of astragalus, calcaneus, and navicular, showing the extreme modification of the hylobatid and the divergence of the astragalar neck. Below: distal view of astragalus and calcaneus. Scale represents 1 cm. For abbreviations see Table 1.

FOOT OF OREOPITHECUS Figure 24. Comparisons, in lateral view, of the articulated astragalus and calcaneus in Hylobates sp. (above) and Pliopithecus (below). Scale represents 1 cm. Note the complete lack of a plantar process and the curiously "bent" proximal end of the calcaneus in Hylobates. In Pliopithecus a heel process (arrow), although not hook shaped or as well developed as in living pongids or Oreopithecus, is clearly present. For abbreviations see Table 1.

6 19

astragalar neck, fitted into a restrictive navicular (Figure 23) is unique and it is likely highly derived among catarrhines. The presence of primitive primate extreme LAJ helical mechanics and the well developed calcaneal plantar process (reduced in gibbons) in Oreopithecus, Pliopithecus, and the Miocene hominoids strongly suggests that the climbing specializations seen in the latter may be ancient indeed. Conclusions From this superficially drawn background the specializations of Oreopithecus are quite puzzling. Oreopithecus does not have to be specially related to the cercopithecids or parapithecids to find a very suitable ancestral source among the Oligocene propliopithecines. The characters of the pes, along with its dental attributes, cannot be readily reconciled with the Miocene hominoid radiation (or at least with its better known members) as subtle suggestions from the morphology of some of the Miocene dryopithecines would align the latter with the living great apes. We can state fairly certainly that the slow climbing adapted foot of Oreopithecuswas extremely chimpanzee-like in function and probably in its overall biorole, but probably evolved independently from the radiation of either the Ponginae or Gorillinae, and possibly independently from the Dryopithecinae. On the other hand one must temper this view with the realization that Oreopithecus shares an elongated ectotympanic with cercopithecids and hominoids (Delson & Szalay, in prep.) and that what can be judged from the morphology of the ulna about the elbow joint suggests great special similarity to hylobatids and gorillines and pongines. If the foot could acquire such chimp-like form-function attributes independently (hinted by the dissimilarity of some important details), then is it so unlikely that the ectotympanic and the humeroulnar articulation evolved convergently also? We have no convincing arguments, except to suggest that the dental morphology and the zygomatic structure of

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Oreopithecus c a n n o t as yet be r e a d i l y reconciled w i t h the M i o c e n e evidence, H a r r i s o n ' s (1986) i n t r i g u i n g few w o r n s p e c i m e n s n o t w i t h s t a n d i n g . T h e M i o c e n e h o m i n o i d s , like g i b b o n s a n d gorillines, t e n d to h a v e a m o d e r a t e l y large t a l a r h e a d a n d short n e c k ( L a n g d o n , 1986). O n e u n a t t r i b u t e d a s t r a g a l u s ( K N M - S O 1705), d i s t i n c t from the o t h e r A f r i c a n s p e c i m e n s , shows r e s e m b l e n c e to Oreopithecus in this region. T h i s s p e c i m e n is the closest m a t c h for Oreopithecus o b s e r v e d a m o n g l i v i n g or fossil c a t a r r h i n e s . W h e t h e r this has a n y p h y l o g e n e t i c significance is yet to be s u p p o r t e d by b e t t e r s p e c i m e n s . W i t h the r e c e n t d i s c o v e r y of w o r n d e n t a l r e m a i n s in E a s t Africa s o m e w h a t r e s e m b l i n g Oreopithecus ( H a r r i s o n , 1986, 1987) the A f r i c a n p r e s e n c e o f o r e o p i t h e c i d s is n o t entirely implausible.

Acknowledgements
W e w o u l d like to express of g r a t i t u d e to D r J o h a n n e s Hfirzeler for a l l o w i n g the s t u d y of s p e c i m e n s in 1972. T h e d r a w i n g s were skillfully p r e p a r e d b y M s P a t r i c i a V a n T a s s e l . S u p p o r t for this project was from the C i t y U n i v e r s i t y of N e w Y o r k D o c t o r a l F a c u l t y R e s e a r c h G r a n t s 01973, 663094 a n d 664296.

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